Description

Upper to lower shore sand- or pebble-scoured cave walls characterized by an impoverished faunal assemblage which may include bryozoan crusts, scattered sponges Halichondria panicea, barnacles such as Semibalanus balanoides or often large Balanus crenatus and the limpet Patella vulgata. The isopod Ligia oceanica may seek refuge in crevices in the rock, and due to the decreased effect of desiccation in these damp caves, other species such as the anemone Actinia equina and spirorbid polychaetes are able to extend further up the shore than normally found on open rock. The lower section of the wall which is subject to greatest scour may be characterized by a band of Spirobranchustriqueter and spirorbid tube-forming polychaetes. In wave sheltered conditions, this biotope may extend to the cave ceiling. The rear of caves on the lower shore may support only sparse fauna consisting of spirorbid polychaetes and barnacles such as Chthamalus montagui with scattered Spirobranchus sp., scattered bryozoan and coralline crusts and in the south-west, occasional Sabellaria alveolata. Shade-tolerant red algae such as Lomentaria articulata may occasionally occur. Due to the low species abundance in this biotope, there may be a variation from cave to cave, depending on local conditions (JNCC, 2015).

Depth range

Mid shore

Additional information

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Further information sources

Sensitivity characteristics of the habitat and relevant characteristic species

The biotope description and characterizing species is taken from JNCC (2015). This biotope is characterized by an impoverished faunal assemblage that typically includes barnacles such as Chthamalus spp., Semibalanus balanoides or Balanus crenatus. Where scour is greatest the biotope may be characterized by a band of Spirobranchus (previously Pomatoceros) triqueter and spirorbid tube-forming polychaetes. As the barnacles and spirorbids are named in the biotope description they are considered key defining species and the sensitivity assessments are focussed on these species. oralline crusts are typically present in this biotope and their sensitivity is assessed. Although other species may be associated with this biotope, such as shade-tolerant red algae, limpets, isopods, anemones and sponges, these are present in low abundances and are not considered to be characterizing and are therefore considered only generally within the assessments.

Abrasion resulting from scouring by sand and pebbles is a key factor structuring the biotope and significant alteration to scouring is likely to change the character of the biotope. Where pressures may alter this factor, this is identified and discussed within the sensitivity assessments.

Resilience and recovery rates of habitat

This biotope is subject to high levels of physical disturbance from abrasion. The species that characterize it are therefore robust animals that can withstand some physical disturbance and/or recover rapidly. This biotope is therefore considered to have a high recovery potential. Sponges and anemones can repair damage and regenerate from small, surviving body parts. Other species such as limpets and ispods are mobile and can migrate into the biotope as adults, while other attached species such as the barnacles and spirorbids produce large numbers of pelagic larvae that can recolonize suitable habitats. Most of the epifauna is probably subject to severe physical disturbance and scour during winter storms and probably develops annually, through regrowth, recolonization and migration from adjacent habitats. Therefore, recovery is likely to be rapid as a typical biological assemblage develops within less than year and probably within 6 months in spring and summer.

Spirorbids are rapid colonisers but poor competitors and hence are maintained in this biotope by the high levels of physical disturbance. Recovery may be within as little as 3 months for these species, based on rapid settlement on artificial panels (Saunders & Connell, 2001; James & Underwood, 1994).Sebens (1985, 1986) observed that calcareous tube worms, encrusting bryozoans and erect hydroids and bryozoans covered scraped areas within 4 months in spring, summer and autumn. Similarly, Hiscock (1983) noted that a community, under conditions of scour and abrasion from stones and boulders moved by storms, developed into a community similar to this biotope, consisting of fast growing species such as Spirobranchus (formerly Pomatoceros) triqueter. Off Chesil Bank, the epifaunal community dominated by Spirobranchus (as Pomatoceros)triqueter and Balanuscrenatus decreased in cover in October as it was scoured away in winter storms, but recolonised in May to June (Gorzula 1977). Warner (1985) reported that the community did not contain any persistent individuals but that recruitment was sufficiently predictable to result in a dynamic stability and a similar community, dominated by Spirobranchus (as Pomatoceros) triqueter, Balanus crenatus and Electra pilosa, (an encrusting bryozoan), was present in 1979, 1980 and 1983 (Riley and Ballerstedt, 2005).

On rocky shores, barnacles are often quick to colonise available gaps, although a range of factors, as outlined below, will influence whether there is a successful episode of recruitment in a year to re-populate a shore following impacts. Bennell (1981) observed that barnacles that were removed when the surface rock was scraped off in a barge accident at Amlwch, North Wales returned to pre-accident levels within 3 years. Petraitis & Dudgeon (2005) also found that Semibalanus balanoides quickly recruited (present a year after and increasing in density) to experimentally cleared areas within the Gulf of Maine, that had previously been dominated by Ascophyllum nodosum However, barnacle densities were fairly low (on average 7.6 % cover) as predation levels in smaller patches were high and heat stress in large areas may have killed a number of individuals (Petraitis et al., 2003). Following creation of a new shore in the Moray Firth, Semibalanus balanoides did not recruit in large numbers until 4 years after shore creation (Terry & Sell, 1986).

Successful recruitment of high number of Semibalanus balanoides individuals to replenish the population may be episodic (Kendall et al., 1985). After settlement the juveniles are subject to high levels of predation as well as dislodgement from waves and sand abrasion depending on the area of settlement. Semibalanus balanoides may live up to 4 years in higher areas of the shore (Wethey,1985). Predation rates are variable (see Petraitis et al., 2003) and are influenced by a number of factors including the presence of algae (that shelters predators such as the dog whelk, Nucella lapillus, and the shore crab, Carcinus maenas and the sizes of clearings (as predation pressure is higher near canopies (Petraitis et al., 2003). Local environmental conditions, including surface roughness (Hills & Thomason, 1998), wind direction (Barnes, 1956), shore height, wave exposure (Bertness et al., 1991) and tidal currents (Leonard et al., 1998) have been identified, among other factors, as factors affecting settlement of Semibalanus balanoides. Biological factors such as larval supply, competition for space, presence of adult barnacles (Prendergast et al., 2009) and the presence of species that facilitate or inhibit settlement (Kendall, et al., 1985, Jenkins et al., 1999) also play a role in recruitment. Mortality of juveniles can be high but highly variable, with up to 90 % of Semibalanus balanoides dying within ten days (Kendall et al., 1985). Presumably these factors also influence the transport, supply and settlement of Chthamaus spp., Balanus crenatus and other species such as spirorbids that produce pelagic larvae.

Resilience assessment. Resilience is assessed as ‘High’ (within 2 years) for all levels of impact, even where resistance is none, as it is likely that a similar community can rapidly develop.

NB: The resilience and the ability to recover from human induced pressures is a combination of the environmental conditions of the site, the frequency (repeated disturbances versus a one off event) and the intensity of the disturbance. Recovery of impacted populations will always be mediated by stochastic events and processes acting over different scales including, but not limited to, local habitat conditions, further impacts and processes such as larval-supply and recruitment between populations. Full recovery is defined as the return to the state of the habitat that existed prior to impact. This does not necessarily mean that every component species has returned to its prior condition, abundance or extent but that the relevant functional components are present and the habitat is structurally and functionally recognisable as the initial habitat of interest. It should be noted that the recovery rates are only indicative of the recovery potential.

Hydrological Pressures

Examples of distribution and thermal tolerances tested in laboratory experiments are provided as evidence to support the sensitivity assessment. In general, populations can acclimate to prevailing conditions which can alter tolerance thresholds and care should, therefore, be used when interpreting reported tolerances. Species that are found in the intertidal are exposed to extremes of high and low air temperatures during periods of emersion. They also experience temperature fluctuation over a short period of time during the tidal cycle. In winter air temperatures are colder than the sea, conversely in summer air temperatures are much warmer than the sea. In general intertidal species are therefore able to tolerate a wide range of temperatures. Within this biotope the cave habitat provides some shade and hence cooler temperatures and reduced desiccation supporting species typically found lower on the shore such as Balanus crenatus and encrusting corallines.

Spirorbids are distributed circumglobally, with some differences in range between taxonomic groups (Knight-Jones et al., 1991). the biotope classification does not specify species or families and therefore it is considered that a spirorbid group may be able to colonize this biotope regardless of temperature changes.

The barnacles Semibalanus balanoides and Balanus crenatus are both ‘northern species’. Semibalanus balanoides extend from Portugal or Northern Spain to the Arctic circle. Populations in the southern part of England are therefore relatively close to the southern edge of their geographic range. Semibalanus balanoides are found on the mid-shore but are less resistant to desiccation that the 'southern' Chthamalus barnacle species. Long-term time series show that successful recruitment of Semibalanus balanoides is correlated to sea temperatures (Mieszkowska, et al., 2014) and that due to recent warming its range has been contracting northwards. Temperatures above 10 to 12 oC inhibit reproduction (Barnes, 1957, 1963, Crisp & Patel, 1969) and laboratory studies suggest that temperatures at or below 10 oC for 4-6 weeks are required in winter for reproduction, although the precise threshold temperatures for reproduction are not clear (Rognstad et al., 2014). Observations of recruitment success in Semibalanus balanoides throughout the South West of England, strongly support the hypothesis that an extended period (4-6 weeks) of sea temperatures <10 oC is required to ensure a good supply of larvae (Rognstad et al., 2014, Jenkins et al., 2000). During periods of high reproductive success, linked to cooler temperatures, the range of barnacles has been observed to increase, with range extensions in the order of 25 km (Wethey et al., 2011), and 100 km (Rognstad et al., 2014).

Balanus crenatus is described as a boreal species (Newman & Ross, 1976) it is found throughout the northeast Atlantic from the Arctic to the west coast of France as far south as Bordeaux; east and west coasts of North America and Japan. In Queens Dock, Swansea where the water was on average 10°C higher than average due to the effects of a condenser effluent, Balanus crenatus was replaced by the subtropical barnacle Balanus amphitrite. After the water temperature cooled Balanus crenatus returned (Naylor, 1965). The increased water temperature in Queens Dock is greater than an increase at the pressure benchmark (2-5°C). Balanus crenatus has a peak rate of cirral beating at 20°C and all spontaneous activity ceases at about 25°C (Southward, 1955). The tolerance of Balanus crenatus, collected in the summer (and thus acclimated to higher temperatures), to increased temperatures was tested in the laboratory. The median upper lethal temperature tolerance was -25.2°C (Davenport & Davenport, 2005) confirming the observations of Southward (1955).

Increased temperatures are likely to favour Chthamalid barnacles present in the biotope rather than Semibalanus balanoides (Southward et al. 1995) and Balanus crenatus. Chthamalus montagui and Chthamalus stellatus are warm water species, with a northern limit of distribution in Britain so are likely to be tolerant of long term increases in temperature. The range of Chthamalus stellatus and Chthamalus montagui has been extending northwards due to increasing temperatures. Chthamalus suffers failure of fertilization at temperatures of 9 °C and below (Patel & Crisp, 1960) , its lower critical temperature for feeding activity is 4.6 °C (Southward, 1955). Semibalanus balanoides outcompetes Chthamalus species for space, but recruitment declines and failures of Semibalanus balanoides in response to warmer temperatures benefit Chthamalus species by allowing them to persist and recruit (Mieszkowska, et al., 2014).

The characterizing Spirobranchus triqueter are found in both warmer and colder waters experienced in the UK. Spirobranchus triqueter occurs from the Arctic, the eastern North Atlantic up to the Mediterranean, Adriatic, Black and Red Sea, the English Channel, the whole North Sea, Skagerrak, Kattegat, the Belts and Öresund up to Bay of Kiel (De Kluijver et al., 2016)

The body temperature of Patella vulgata can exceed 36oC in the field, (Davies, 1970), adults become become non-responsive at 37-38oC and die at temperatures of 42oC (Evans, 1948). Lower temperatures enhance feeding rates in adults (Thompson et al., 2004). Juvenile tolerance of warm air temperatures and dessication may be lower than adults. Juveniles require damp areas of rock (Lewis & Bowmna, 1975) and the bare rock surfaces typical of this biotope, present a harsher habitat than the associated crevices and cracks. Long-term time studies in southern England suggest that Patella vulgata have become scarcer following warmer summers, while Patella depressa increase in abundance (Southward et al., 1995). Increased temperatures may alter spawning cues and reproduction success in Patella vulgata populations. Observations suggest that spawning is initiated in autumn storms with greater wave action when seawater temperatures drop below 12oC (Bowman 1985, Bowman & Lewis, 1986, LeQuesne, 2005). In Northern Portugal warming seas appear to be linked to both a shortening of the reproductive period and the lack of multiple spawning events in Patella vulgata and other northern species (Ribeiro et al., 2009).

The encrusting coralline, Lithophyllum incrustans, is close to the northern edge of its reported distribution range in the UK (Kain, 1982; Guiry & Guiry, 2015) and is therefore considered likely to be tolerant of an increase in temperature, particularly in this subtidal biotope, where it is protected from desiccation.

Sensitivity assessment. Typical surface water temperatures around the UK coast vary, seasonally from 4-19°C (Huthnance, 2010). The biotope is considered to tolerate a 2°C increase in temperature for a year. An acute increase at the pressure benchmark may be tolerated in winter, but a sudden return to typical temperatures could lead to mortalities among acclimated animals. No evidence was found to support this assessment, however. An acute increase of 5°C in summer would be close to the lethal thermal temperature for Balanus crenatus. Biotope resistance is therefore assessed as ‘Medium’ and resilience as ‘High’ and biotope sensitivity is therefore ‘Low’. Adult Semibalanus balanoides are considered likely to be able to tolerate an acute or chronic change, however, if an acute change in temperature occurred in autumn or winter it could disrupt reproduction, while a chronic change could alter reproductive success if it exceeded thermal thresholds for reproduction. The effects would depend on the magnitude, duration and footprint of the activities leading to this pressure. However, barnacle populations are highly connected, with a good larval supply and high dispersal potential (Wethey et al., 2011, Rognstad et al., 2014). Resistance is therefore assessed as ‘High’ (despite some potential effects on reproductive success) and resilience as ‘High’ (by default). This biotope is therefore considered to be ‘Not sensitive’ at the pressure benchmark, although some changes in the proportions of different barnacle species may occur.

Many intertidal species are tolerant of freezing conditions as they are exposed to extremes of low air temperatures during periods of emersion. They must also be able to cope with sharp temperature fluctuations over a short period of time during the tidal cycle. In winter air temperatures are colder than the sea, conversely in summer air temperatures are much warmer than the sea. Species that occur in the intertidal are therefore generally adapted to tolerate a range of temperatures, with the width of the thermal niche positively correlated with the height of the shore that the animal usually occurs at (Davenport & Davenport, 2005).

Spirorbids are distributed circumglobally, with some differences in range between taxonomic groups (Knight-Jones et al., 1991). the biotope classification does not specify species or families and therefore it is considered that a spirorbid group may be able to colonize this biotope regardless of temperature changes.

The barnacle Semibalanus balanoides is primarily a ‘northern’ species with an arctic-boreal distribution. Long-term time series show that recruitment success is correlated to lower sea temperatures (Mieszkowska et al., 2014). Due to warming temperatures its range has been contracting northwards. Temperatures above 10 to 12 oC inhibit reproduction (Barnes, 1957, 1963, Crisp & Patel, 1969) and laboratory studies suggest that temperatures at or below 10 oC for 4-6 weeks are required in winter for reproduction, although the precise threshold temperatures for reproduction are not clear (Rognstad et al., 2014). The tolerance of Semibalanus balanoides collected in the winter (and thus acclimated to lower temperatures) to low temperatures was tested in the laboratory. The median lower lethal temperature tolerance was -14.6 oC (Davenport & Davenport, 2005). A decrease in temperature at the pressure benchmark is therefore unlikely to negatively affect this species. Balanus crenatus is described as a boreal species (Newman & Ross, 1976) it is found throughout the northeast Atlantic from the Arctic to the west coast of France as far south as Bordeaux; east and west coasts of North America and Japan. Chthamalus stellatus and Chthamalus montagui are ‘southern’ barnacle species and their range has been extending northwards due to increasing temperatures. Chthamalus suffers failure of fertilization at temperatures of 9 °C and below (Patel and Crisp, 1960) its lower critical temperature for feeding activity is 4.6 °C (Southward, 1955). The cold winter of 2009-10 in France led to recruitment failure in Chthamalus species (Wethey et al., 2011).

The characterizing Spirobranchus triqueter are found in both warmer and colder waters experienced in the UK. Spirobranchus triqueter occurs from the Arctic, the eastern North Atlantic up to the Mediterranean, Adriatic, Black and Red Sea, the English Channel, the whole North Sea, Skagerrak, Kattegat, the Belts and Öresund up to Bay of Kiel (De Kluijver et al., 2016)

The limpet, Patella vulgata is largely unaffected by short periods of extreme cold. Ekaratne & Crisp (1984) found adult limpets continuing to grow over winter when temperatures fell to -6°C, and stopped only by still more severe weather. However, loss of adhesion after exposure to -13°C has been observed with limpets falling off rocks and therefore becoming easy prey to crabs or birds (Fretter & Graham, 1994). However, in the very cold winter of 1962-3 when temperatures repeatedly fell below 0°C over a period of 2 months large numbers of Patella vulgata were found dead (Crisp, 1964). Periods of frost may also kill juvenile Patella vulgata, resulting in recruitment failures in some years (Bowman & Lewis, 1977).

Sensitivity assessment. Based on the wide temperature tolerance range of the characterizing and associated species it is concluded that the acute and chronic decreases in temperature described by the benchmark would have limited effect on barnacles and limpets. Based on distribution and the link between cooler winter temperatures and reproductive success, Semibalanus balanoides is also considered to be unaffected at the pressure benchmark. Changes in recruitment success between southern and northern barnacle species may lead to some changes in the proportions of barnacles species recruiting but this is not considered to alter the character of the biotope. Resistance is therefore assessed as ‘High’ (despite some potential effects on reproductive success) and resilience as ‘High’ (by default). This biotope is therefore considered to be ‘Not sensitive’ at the pressure benchmark

This biotope is recorded in full salinity (30-35 ppt) habitats (Connor et al., 2004) and therefore the sensitivity assessment benchmark considers an increase from full salinity to >40 ppt. Biotopes found in the intertidal will naturally experience fluctuations in salinity where evaporation increases salinity and inputs of rainwater expose individuals to fresh water. Species found in the intertidal are therefore likely to have some form of behavioural or physiological adaptations to changes in salinity.

Barnes & Barnes (1974) found that larvae from six barnacle species including Balanus crenatus, Chthamalus stellatus and Semibalanus (as Balanus) balanoides, completed their development to nauplii larvae at salinities between 20-40‰. (Some embryos exposed at later development stages could survive at higher and lower salinities). Balanus crenatus occurs in estuarine areas and is therefore adapted to variable salinity (Davenport, 1976). When subjected to sudden changes in salinity Balanus crenatus closes its opercular valves so that the blood is maintained temporarily at a constant osmotic concentration (Davenport, 1976).

The crustose corallines that occur in this biotope may also be found on rocky shores and in rockpools where salinities may fluctuate markedly during exposure to the air. Edyvean & Ford (1984b) suggest that populations of Lithophyllum incrustans are affected by temperature changes and salinity and that temperature and salinity ‘shocks’ induce spawning but no information on thresholds was provided (Edyvean & Ford, 1984b). Populations of Lithophyllum incrustans were less stable in rockpools with a smaller volume of water that were more exposed to temperature and salinity changes due to lower buffering capacity. Sexual plants (or the spores that give rise to them) were suggested to be more susceptible than asexual plants to extremes of local environmental variables (temperature, salinity, etc.) as they occur with greater frequency at sites where temperature and salinity were more stable (Edyvean & Ford, 1984b).

Local populations may be acclimated to the prevailing salinity regime and may therefore exhibit different tolerances to other populations subject to different salinity conditions and therefore caution should be used when inferring tolerances from populations in different regions. This biotope is found in full (30-35 ppt) salinity (Connor et al., 2004). Biotopes found in the intertidal will naturally experience fluctuations in salinity where evaporation increases salinity and inputs of rainwater expose individuals to fresh water. Species found in the intertidal are therefore likely to have some form of behavioural or physiological adaptations to changes in salinity.

Sensitivity assessment. Little direct evidence was found to assess sensitivity to this pressure. Although some increases in salinity may be tolerated by the associated species present these are generally short-term and mitigated during tidal inundation. This biotope is considered, based on distribution on the mid-shore to be sensitive to a persistent increase in salinity to > 40 ppt. Resistance is therefore assessed as ‘Low’ and recovery as ‘High’ (following restoration of usual salinity). Sensitivity is therefore assessed as ‘Low'.

This biotope is recorded in full salinity (30-35 ppt) (Connor et al., 2004). At the pressure benchmark, a change from full to variable salinity (18-30 ppt) is assessed. The characterizing species Balanus crenatus and Spirobranchus triqueter are found in a similar biotope (CR.MCR.EcCr.UrtScr), is present in variable salinities (Connor et al., 2004). It is therefore likely that the characterizing species will tolerate a reduction in salinity form full to reduced. No direct evidence was found to assess the sensitivity of spirorbids, however, these are described as frequent in the variable salinity biotope LR.LLR.FVS.FserVSIR. and the low salinity biotope LIR.Lag.ProtFur. It is therefore inferred that this group would not be impacted by a reduction in salinity at the pressure benchmark.

Edyvean & Ford (1984b) suggest that populations of the crustose coralline Lithophyllum incrustans are affected by temperature changes and salinity and that temperature and salinity ‘shocks’ induce spawning but no information on thresholds were provided (Edyvean & Ford, 1984b). Populations of Lithophyllum incrustans were less stable in tide pools with a smaller volume of water that were more exposed to temperature and salinity changes due to lower buffering capacity. Sexual plants (or the spores that give rise to them) were suggested to be more susceptible than asexual plants to extremes of local environmental variables (temperature, salinity etc.) as they occur with greater frequency at sites where temperature and salinity were more stable (Edyvean & Ford, 1984b).

Balanus crenatus occurs in estuarine areas and is therefore adapted to variable salinity (Davenport, 1976). When subjected to sudden changes in salinity Balanus crenatus closes its opercular valves so that the blood is maintained temporarily at a constant osmotic concentration (Davenport, 1976). Acclimation to different salinity regimes alters the point at which opercular closure and resumption of activity occurs (Davenport, 1976). Balanus crenatus can tolerate salinities down to 14 psu if given time to acclimate (Foster, 1970). At salinities below 6 psu motor activity ceases, respiration falls and the animal falls in to a "salt sleep". In this state the animals may survive (Barnes & Barnes, 1974) in fresh water for 3 weeks, enabling them to withstand changes in salinity over moderately long periods (Barnes & Powell, 1953). Larvae are more sensitive than adults. In culture experiments, eggs maintained below 10‰ rupture, due to osmotic stress (Barnes & Barnes, 1974). At 15-17‰ there is either no development of early stages or the nauplii larvae are deformed and “probably not viable”, similarly at 20‰ development occurs, but about half of the larvae are deformed and not viable. (Barnes & Barnes, 1974). Normal development resulting in viable larvae occurs between salinities of 25-40 ‰ (Barnes & Barnes, 1974). Barnes & Barnes (1965) found that in high suspended solids and low salinity there was a decrease in the number of eggs per brood of Chthamalus stellatus / Chthamalus montagui If salinities decrease below 21 psu all cirral activity of barnacles normally associated with full salinity waters, ceases (Foster, 1971).Semibalanus balanoides are tolerant of a wide range of salinity and can survive periodic emersion in freshwater, e.g. from rainfall or fresh water run off, by closing their opercular valves (Foster, 1971b). They can also withstand large changes in salinity over moderately long periods of time by falling into a "salt sleep" and can be found on shores (example from Sweden) with large fluctuations in salinity around a mean of 24 (Jenkins et al., 2001).

Spirobranchus triqueter has not been recorded from brackish or estuarine waters. Therefore, it is likely that the species will be very intolerant of a decrease in salinity. However, Dixon (1985, cited in Riley & Ballerstedt, 2005) views the species as able to withstand significant reductions in salinity. The degree of reduction in salinity and time that the species could tolerate those levels were not recorded. Therefore, there is insufficient information available to assess the intolerance of Spirobranchus triqueter to a reduction in salinity and the assessment is based on its presence in the biotope CR.MCR.EcCr.UrtScr which occurs in variable salinity (as well as full) habitats (Connor et al., 2004).

Sensitivity assessment. As the characterizing species are found in biotopes in both full and variable salinity habitats, the biotope is considered ‘Not sensitive’ to a decrease in salinity from full to variable. Biotope resistance is therefore assessed as ‘High’ and resilience is assessed as ‘High’ (by default) and the biotope is assessed as ‘Not sensitive’.

The spirorbids, barnacles and coralline crusts characterizing this biotope are securely attached and as these are relatively flat and small they are subject to little or no drag compared to upright growth forms. Colonies of Lithophyllum incrustans appear to thrive in conditions exposed to strong water movement (Irvine & Chamberlain, 1994

Changes in flow rate may impact the supply of food to filter feeders. Laboratory experiments demonstrate that barnacle feeding behaviour alters over different flow rates but that barnacles can feed at a variety of flow speeds (Sanford et al., 1994). Flow tank experiments using velocities of 0.03, 0.07 and 0.2 m/s showed that a higher proportion of barnacles fed at higher flow rates (Sanford et al., 1994). Feeding was passive, meaning the cirri were held out to the flow to catch particles; although active beating of the cirri to gernerate feeding currents occurs in still water (Crisp & Southward, 1961). Field observations at sites in southern New England (USA) that experience a number of different measured flow speeds, found that Semibalanus balanoides from all sites responded quickly to higher flow speeds, with a higher proprtion of individuals feeding when current speeds were higher. Barnacles were present at a range of sites, varying from sheltered sites with lower flow rates (maximum observed flow rates <0.06- 0.1 m/s), a bay site with higher flow rates (maximum observed flows 0.2-0.3 m/s) and open coast sites (maximum observed flows 0.2-0.4 m/s). Recruitment was higher at the site with flow rates of 0.2-0.3 m/s (although this may be infuenced by supply) and at higher flow microhabitats within all sites. Both laboratory and field observations indicate that flow is an important factor with effects on feeding, growth and recruitment in Semibalanus balanoides (Sanford et al., 1994, Leonard et al., 1998), however, the results suggest that flow is not a limiting factor determining the overall distribution of barnacles as they can adapt to a variety of flow speeds.

Spirobranchus triqueter is found in biotopes exposed to flow speeds varying from very weak to moderately strong (negligible - >1.5m/s) and was considered ‘Not sensitive’ at the pressure benchmark (Tillin & Tyler-Walters, 2014). Balanus crenatus is found in a very wide range of water flows (Tillin & Tyler-Walters, 2014), although it usually occurs in sites sheltered from wave action (Eckman & Duggins, 1993) and can adapt feeding behaviour according to flow rates. In the absence of any current, the barnacle rhythmically beats its cirri to create a current to collect zooplankton. Growth of Balanus crenatus (measured as increase in basal area), maintained for 69 days at constant flow speeds in laboratory experiments was greatest at intermediate flow speeds (0.08 m/s) and decreased at higher speeds (Eckman & Duggins, 1993). Over the entire range of flow speeds measured (0.02 m/s – 0.25 m/s), Balanus crenatus, was able to control the cirrus with little or no deformation by flow observed (Eckman, & Duggins, 1993).

Sensitivity assessment. The species that characterize or are associated with this biotope are securely attached and can occur in a range of flow speeds. Resistance of the biotope to changes in water flow is assessed as ‘High’ and resilience as ‘High’ (by default) so that the biotope is assessed as ‘Not sensitive’. Scour is a key factor structuring this biotope (Connor et al., 2004), changes in flow exceeding the pressure benchmark may increase or decrease sediment transport and associated scour may lead to indirect changes in the character of the biotope.

Emergence regime is a key factor structuring this (and other) intertidal biotopes. Increased emergence may reduce habitat suitability for characterizing species through greater exposure to dessication and reduced feeding opportunities for the barnacles and other filter feeders including spirobids, barnacles sponges and anemones which feed when immersed. Semibalanus balanoides is less tolerant of dessication stress than Chthamalus barnacles species and changes in emergence may therefore lead to species replacement and the development of a Chthamalus sp. dominated biotope, more typical of the upper shore may develop. Records suggest that, typically, above this biotope the community may grade into a red algal dominated community (AudCla), where light allows (Connor et al., 2004). Changes in emergence may therefore eventually lead to replacement of this biotope to one more typical of the upper cave walls.

Decreased emergence would reduce dessication stress and allow the attached suspension feeders more feeding time. Predation pressure on barnacles and limpets is likely to increase where these are submerged for longer periods and to prevent colonisation of lower zones. Semibalanus balanoides was able to extend its range into lower zones when protected from predation by the dogwhelk, Nucella lapillus (Connell, 1961). The mobile species present within the biotope, including Patella vulgata, isopods and Littorina spp. would be able to relocate to preferred shore levels. Where decreased emergence leads to increased abrasion and scour while immersed, the removal of fauna may lead to this biotope reverting to the more barren LR.FLR.CvOv.BarCv.

Sensitivity assessment. As emergence is a key factor structuring the distribution of animals on the shore, resistance to a change in emergence (increase or decrease) is assessed as ‘Low’. Recovery is assessed as ‘High’, and sensitivity is therefore assessed as 'Low'.

This biotope is recorded from locations that are judged to range from very exposed to sheltered (JNCC, 2015).

The barnacles, spirorbids and crustose corallines associated with this biotope have a flat growth form and are unlikely to be dislodged by increased wave action.

Sensitivity assessment. The biotope and characterizing and associated species are found across a range of wave exposures, populations occurring within the middle of the range are considered to have 'High' resistance to a change in significant wave height at the pressure benchmark. Resilience is assessed as ‘High’, by default, and the biotope is considered ‘Not sensitive’.

Chemical Pressures

Not sensitive at the pressure benchmark that assumes compliance with all relevant environmental protection standards.

No information was found concerning the effects of heavy metals on encrusting coralline algae. Bryan (1984) suggested that the general order for heavy metal toxicity in seaweeds is: organic Hg> inorganic Hg > Cu > Ag > Zn> Cd> Pb. Contamination at levels greater than the pressure benchmark may adversely impact the biotope. Cole et al. (1999) reported that Hg was very toxic to macrophytes. The sub-lethal effects of Hg (organic and inorganic) on the sporelings of an intertidal red algae, Plumaria elegans, were reported by Boney (1971). 100% growth inhibition was caused by 1 ppm Hg.

Contamination at levels greater than the pressure benchmark may adversely impact the biotope. Barnacles accumulate heavy metals and store them as insoluble granules (Rainbow, 1987). Pyefinch & Mott (1948) recorded a median lethal concentration of 0.19 mg/l copper and 1.35 mg/l mercury, for Balanus crenatus over 24 hours. Barnacles may tolerate fairly high level of heavy metals in nature, for example they are found in Dulas Bay, Anglesey; where copper reaches concentrations of 24.5 µg/l, due to acid mine waste (Foster et al., 1978).

Bryan (1984) suggested that gastropods are also rather tolerant of heavy metals. In the Fal estuary Patella vulgata occurs at, or just outside, Restronguet Point at the end of the creek where metal concentrations are in the order: Zinc (Zn) 100-2000µg/l, copper (Cu) 10-100µg/l and cadmium (Cd) 0.25-5µg/l (Bryan & Gibbs, 1983). However, in the laboratory Patella vulgata was found to be intolerant of small changes in environmental concentrations of Cd and Zn by Davies (1992). At concentrations of 10µg/l pedal mucus production and levels of activity were both reduced, indicating a physiological response to metal concentrations. Exposure to Cu at a concentration of 100µg/l for one week resulted in progressive brachycardia (slowing of the heart beat) and the death of limpets. Zn at a concentration of 5500µg/l produced the same effect (Marchan et al.,1999).

Not sensitive at the pressure benchmark that assumes compliance with all relevant environmental protection standards

Where exposed to direct contact with fresh hydrocarbons, encrusting coralline algae appears to be highly intolerant. Crump et al. (1999) described 'dramatic and extensive bleaching' of 'Lithothamnia' following the Sea Empress oil spill. Observations following the Don Marika oil spill (K. Hiscock, pers. comm.) were of rockpools with completely bleached coralline algae. However, Chamberlain (1996) observed that although Lithophyllum incrustans was affected in a short period of time by oil during the Sea Empress spill, recovery occurred within about a year. The oil was found to have destroyed about one third of the thallus thickness but regeneration occurred from thallus filaments below the damaged area.

No information is available on the intolerance of Balanus crenatus to hydrocarbons. However, other littoral barnacles generally have a high tolerance to oil (Holt et al., 1995) and were little impacted by the Torrey Canyon oil spill (Smith, 1968), so Balanus crenatus is probably fairly resistant to oil.

Not sensitive at the pressure benchmark that assumes compliance with all relevant environmental protection standards.

Barnacles have a low resilience to chemicals such as dispersants, dependant on the concentration and type of chemical involved (Holt et al., 1995). They are less intolerant than some species (e.g. Patella vulgata) to dispersants (Southward & Southward, 1978) and Balanus crenatus was the dominant species on pier pilings at a site subject to urban sewage pollution (Jakola & Gulliksen, 1987). Hoare & Hiscock (1974) found that Balanus crenatus survived near to an acidified halogenated effluent discharge where many other species were killed, suggesting a high tolerance to chemical contamination. Little information is available on the impact of endocrine disrupters on adult barnacles. Holt et al. (1995) concluded that barnacles are fairly sensitive to chemical pollution, therefore intolerance is reported as high. The species is an important early colonizer of sublittoral rock surfaces (Kitching, 1937) and it heavily recolonized a site that was dredged for gravel within 7 months (Kenny & Rees, 1994). Therefore, recovery is predicted to be high.

Hoare & Hiscock (1974) reported that the limpet Patella vulgata was excluded from sites within 100-150m of the discharge of acidified, halogenated effluent in Amlwch Bay. Limpets are also extremely intolerant of aromatic solvent based dispersants used in oil spill clean-up. During the clean-up response to the Torrey Canyon oil spill nearly all the limpets were killed in areas close to dispersant spraying. Viscous oil will not be readily drawn in under the edge of the shell by ciliary currents in the mantle cavity, whereas detergent, alone or diluted in seawater, would creep in much more readily and be liable to kill the limpet (Smith, 1968). A concentration of 5ppm killed half the limpets tested in 24 hours (Southward & Southward, 1978; Hawkins & Southward, 1992). Acidified seawater affects the motility of Patella vulgata. At a pH of 5.5 motility was reduced whilst submerged but individuals recovered when returned to normal seawater. At a pH of 2.5 total inhibition of movement occurred and when returned to normal seawater half had died (Bonner et al., 1993). Reduced motility reduces time for foraging and may result in decreased survival of individuals. Acidified seawater can also change the shell composition which will lead to a decrease in its protective nature and hence survival (Bonner et al., 1993). Short periods (48 hours) are unlikely to have much effect on a population but long periods (1 year) may cause reduced grazing and an increase in algal growth. However, seawater is unlikely to reach pH 2.5 therefore intolerance to slight changes in pH will be low. Hoare & Hiscock (1974) reported that in Amlwch Bay Patella vulgata was excluded from sites within 100-150m of the discharge of acidified, halogenated effluent.

Most pesticides and herbicides were suggested to be very toxic for invertebrates, especially crustaceans (amphipods isopods, mysids, shrimp and crabs) and fish (Cole et al., 1999).

Specific information concerning oxygen consumption and reduced oxygen tolerances were not found for the key characterizing species within the biotope. It is likely that as this biotope occurs in areas that are shallow and tidally flushed that re-oxygenation is likely, limiting the effects of any de-oxygenation events. However, this may mean that the species present have little exposure to low oxygen and may be sensitive to this pressure. Balanus crenatus, however, respires anaerobically so it can withstand some decrease in oxygen levels. When placed in wet nitrogen, where oxygen stress is maximal and desiccation stress is minimal, Balanus crenatus has a mean survival time of 3.2 days (Barnes et al., 1963) and this species is considered to be ‘Not sensitive’ to this pressure. Semibalanus balanoides can respire anaerobically, so they can tolerate some reduction in oxygen concentration (Newell, 1979). When placed in wet nitrogen, where oxygen stress is maximal and desiccation stress is low, Semibalanus balanoides have a mean survival time of 5 days (Barnes et al., 1963).

In laboratory experiments a reduction in the oxygen tension of seawater from 148mm Hg (air saturated seawater) to 50mm Hg rapidly resulted in reduced heart rate in limpets of the genus Patella (Marshall & McQuaid, 1993). Heartbeat rate returned to normal in oxygenated water within two hours. Limpets can survive for a short time in anoxic seawater; Grenon & Walker, (1981) found that in oxygen free water limpets could survive up to 36 hours, although Marshall & McQuaid (1989) found a lower tolerance for Patella granularis, which survived up to 11 hours in anoxic water. Therefore, some individuals may survive for one week at an oxygen concentration of 2mg/l. However, Patella vulgata is able to respire in air, so would only be exposed to low oxygen in the water column intermittently during periods of tidal immersion. In addition, in areas of wave exposure and moderately strong current flow low oxygen levels in the water are unlikely to persist for very long.

Sensitivity assessment. Based on Balanus crenatus and Semibalanus balanoides and mitigation of de-oxygenation by water movements, this biotope is considered to have 'High' resistance and High resilience (by default), and is therefore 'Not sensitive'.

Nutrient enrichment at the pressure benchmark is unlikely to affect the fauna within this biotope. No direct evidence was found to assess this pressure. A slight increase in nutrient levels could be beneficial for barnacles by promoting the growth of phytoplankton levels and therefore increasing zooplankton levels. Balanus crenatus was the dominant species on pier pilings, which were subject to urban pollution (Jakola & Gulliksen, 1987). Limpets and other grazers would also benefit from increased growth of benthic microalgae. However, Holt et al. (1995) predict that smothering of barnacles by ephemeral green algae is a possibility under eutrophic conditions.

Over geological timescales, periods of increased nutrient availability have experienced increases in the distribution of crustose coralline species at the expense of corals (Littler & Littler, 2013), suggesting that this group have some tolerance for enhanced nutrient levels. Overall, Littler & Littler (2013) suggest that corallines as a group can tolerate both low and elevated levels of nutrients. The encrusting coralline Lithophyllum incrustans were present at sites dominated by Ulva spp. in the Mediterranean exposed to high levels of nutrient enrichment from domestic sewage (Arévalo et al., 2007).

Sensitivity assessment. The pressure benchmark is set at a level that is relatively protective and based on the evidence and considerations outlined above the biological assemblage is considered to be 'Not sensitive' at the pressure benchmark. Resistance and resilience are therefore assessed as 'High'.

As the biotope occurs in tide swept or wave exposed areas (Connor et al., 2004), water movements will disperse organic matter reducing the level of exposure.

The animals found within the biotope may be able to utilise the input of organic matter as food, or are likely to be tolerant of inputs at the benchmark level. Cabral-Oliveira et al., (2014), found that filter feeders including the barnacle Chthamalus montagui, were more abundant at sites closer to a sewage treatment works, as they could utilise the organic matter inputs as food. On the same shores, higher abundances of juvenile Patella sp. and lower abundances of adults were found closer to sewage inputs, Cabral-Oliveira et al., (2014) suggested the structure of these populations was due to increased competition closer to the sewage outfalls.

In a recent review, assigning species to ecological groups based on tolerances to organic pollution, characterizing animal species; Balanus crenatus and Spirobranchus triqueter were assigned to AMBI Group II described as 'species indifferent to enrichment, always present in low densities with non-significant variations with time, from initial state, to slight unbalance' (Gittenberger & Van Loon, 2011).

The crusting coralline Lithophyllum incrustans were present at sites dominated by Ulva spp. in the Mediterranean exposed to high levels of organic pollution from domestic sewage (Arévalo et al., 2007), suggesting the encrusting corallines are not sensitive to this pressure.

Sensitivity assessment. It is not clear whether the pressure benchmark would lead to enrichment effects in this dynamic, scoured habitat. High water movements would disperse organic matter particles, mitigating the effect of this pressure. Based on the AMBI categorisation (Borja et al., 2000, Gittenberger & Van Loon, 2011), characterizing and associated species are assessed as ‘Not Sensitive’ to this pressure based on ‘High’ resistance and ‘High’ resilience as there is no impact to recover from. Although species within the biotope may be sensitive to gross organic pollution resulting from sewage disposal and aquaculture they are considered to have ‘High’ resistance to the pressure benchmark (which represents organic enrichment) and therefore ‘High’ resilience. The biotope is therefore considered to be ‘Not Sensitive’.

Physical Pressures

All marine habitats and benthic species are considered to have a resistance of ‘None’ to this pressure and to be unable to recover from a permanent loss of habitat (resilience is ‘Very Low’). Sensitivity within the direct spatial footprint of this pressure is therefore ‘High’. Although no specific evidence is described confidence in this assessment is ‘High’, due to the incontrovertible nature of this pressure.

This biotope is characterized by the hard rock substratum to which the key characterizing species, spirobids, barnacles and associated species can firmly attach to. A change to a sedimentary substratum would significantly alter the character of the biotope. More subtle changes in substratum type can also lead to indirect effects. Surface roughness, for example, is correlated with settlement in barnacles (Coombes et al., 2015). Spirorbids are also selective and will discriminate between different types of hard surface (James & Underwood, 1994). An artificial substratum may therefore not be equivalent to a natural rock reef habitat. An increase in mobile surfaces can also indirectly decrease suitable habitats. Shanks & Wright (1986) observed that limpet mortalities were much higher at sites where the supply of loose cobbles and pebbles were greater, leading to increased abrasion through wave action 'throwing' rocks onto surfaces. The biotope is therefore considered to have 'No' resistance to this pressure (based on a change to sediments), recovery is assessed as 'Very low', as the change at the pressure benchmark is permanenet. Biotope sensitivity is therefore assessed as 'High'. As this biotope is found in caves, a change in topography from a cave to an open rock surface would also result in the loss of the biotope.

The species characterizing this biotope are epifauna or epiflora occurring on rock and would be sensitive to the removal of the habitat. However, extraction of rock substratum is considered unlikely and this pressure is considered to be ‘Not relevant’ to hard substratum habitats.

The species characterizing this biotope occur on the rock and therefore have no protection from surface abrasion. High levels of abrasion from scouring by mobile sands and gravels is an important structuring factor in this biotope (Connor et al., 2004) and prevents replacement by less scour-tolerant species, such as shade-tolerant red algae.

The effects of trampling (a source of abrasion) on barnacles appears to be variable with some studies not detecting significant differences between trampled and controlled areas (Tyler-Walters & Arnold, 2008). However, this variability may be related to differences in trampling intensities and abundance of populations studied. The worst case incidence was reported by Brosnan and Crumrine (1994) who reported that a trampling pressure of 250 steps in a 20x20 cm plot one day a month for a period of a year significantly reduced barnacle cover at two study sites. Barnacle cover reduced from 66% to 7% cover in 4 months at one site and from 21% to 5% within 6 months at the second site. Overall barnacles were crushed and removed by trampling. Barnacle cover remained low until recruitment the following spring. Long et al. (2011) also found that heavy trampling (70 humans km-1 shoreline h-1) led to reductions in barnacle cover. Single step experiments provide a clearer, quantitative indication of sensitivity to direct abrasion. Povey & Keough (1991) in experiments on shores in Mornington peninsula, Victora, Australia, found that in single step experiments 10 out of 67 barnacles, (Chthamlus antennatus about 3mm long), were crushed. However, on the same shore, the authors found that limpets may be relatively more resistant to abrasion from trampling. Following step and kicking experiments, few individuals of the limpet Cellana trasomerica, (similar size to Patella vulgata) suffered damage or relocated (Povey & Keough, 1991). One kicked limpet (out of 80) was broken and 2 (out of 80) limpets that were stepped on could not be relocated the following day (Povey & Keough, 1991). Trampling may lead to indirect effects on limpet populations, Bertocci et al., (2011) found that the effects of trampling on Patella sp. increased temporal and spatial variability of in abundance. The experimental plots were sited on a wave-sheltered shore dominated by Ascophyllum nodosum. On these types of shore, trampling in small patches, that removes macroalgae and turfs, will indirectly enhance habitat suitability for limpets by creating patches of exposed rock for grazing.

Mechanical abrasion from scuba divers was reported to impact encrusting corallines, with cover of Lithophyllum stictaeforme greater in areas where diving was forbidden than visited areas (abundance, 6.36 vs 1.4; it is presumed this refers to proportion of cover, although this is not clear from the text, Guarinieri et al., 2012). Dethier (1994) experimentally manipulated surface abrasion on a range of encrusting algae including Lithophyllum impressum. Crusts were brushed with either a nylon or steel brush for 1 minute each month for 24 months. Unbrushed controls grew by approximately 50% where the cover of nylon brushed crusts and steel brushed crusts decreased by approximately 25% and 40% respectively (interpreted from figures in Dethier, 1994). In laboratory tests on chips of Lithophyllum impressum brushing with a steel brush for 1 minute once a week for 3 weeks, resulted in no cover loss of two samples while a third ‘thinned and declined’ (Dethier, 1994).

Hiscock (1983) noted that a community, under conditions of scour and abrasion from stones and boulders moved by storms, developed into a community similar to this biotope, consisting of fast growing species such as Spirobranchus (formerly Pomatoceros) triqueter. Off Chesil Bank, the epifaunal community dominated by Spirobranchus (as Pomatoceros)triqueter and Balanuscrenatus decreased in cover in October as it was scoured away in winter storms, but recolonised in May to June (Gorzula 1977). Warner (1985) reported that the community did not contain any persistent individuals but that recruitment was sufficiently predictable to result in a dynamic stability and a similar community, dominated by Spirobranchus (as Pomatoceros) triqueter, Balanus crenatus and Electra pilosa, (an encrusting bryozoan), was present in 1979, 1980 and 1983 (Riley and Ballerstedt, 2005).

Shanks & Wright (1986), found that even small pebbles (<6 cm) that were thrown by wave action in Southern California shores could create patches in Chthamalus fissus aggregations and could smash owl limpets (Lottia gigantea). Average, estimated survivorship of limpets at a wave exposed site, with many loose cobbles and pebbles allowing greater levels of abrasion was 40% lower than at a sheltered site. Severe storms were observed to lead to almost total destruction of local populations of limpets through abrasion by large rocks and boulders.

Sensitivity assessment. The impact of surface abrasion will depend on the footprint, duration and magnitude of the pressure. High levels of abrasion from scouring by mobile cobbles and pebbles is an important structuring factor in this biotope (Connor et al., 2004) but the persistence of the assemblage may depend on rapid recovery rather than high resistance (Gorzula, 1977). The evidence for the effects of trampling on barnacles and severe scour on Balanus crenatusandSpirobranchus triqueter, suggest that resistance, to a single abrasion event is ‘Low’ and recovery is ‘High’. Sensitivity is assessed as ‘Low’, based upon the information for these species and the Coralline spp.

The species characterizing this biotope group are epifauna or epiflora occurring on rock which is resistant to subsurface penetration. The assessment for abrasion at the surface only is therefore considered to equally represent sensitivity to this pressure.

This biotope occurs in scoured habitats and it is likely, depending on local sediment supply, that the biotope is exposed to chronic or intermittent episodes of high-levels of suspended solids as local sediments are re-mobilised and transported. A significant increase in suspended solids may result in smothering (see siltation pressures) where these are deposited. Based on Cole et al. (1999) and Devlin et al. (2008) this biotope is considered to experience intermediate turbidity (10-100 mg/l) based on UK TAG (2014). An increase at the pressure benchmark refers to a change to medium turbidity (100-300 mg/l) and a decrease is assessed as a change to clear (<10 mg/l) based on UK TAG (2014).

An increase in turbidity could be beneficial if the suspended particles are composed of organic matter, however high levels of suspended solids with increased inorganic particles may reduce filter feeding efficiencies. A reduction in suspended solids will reduce food availability for filter feeding species in the biotope (where the solids are organic), although effects are not likely to be lethal over the course of a year. A reduction in light penetration could also reduce growth rate of phytoplankton and so limit zooplankton levels. However, light penetration itself is unlikely to be an important factor as both Balanus crenatus and Spirobranchus triqueter are recorded from the lower eulittoral or the lower circalittoral.

Available evidence indicates that Spirobranchus triqueter is tolerant of a wide range of suspended sediment concentrations (Riley and Ballerstedt, 2005). Stubbings and Houghton (1964) recorded Spirobranchus (as Pomatoceros) triqueter in Chichester harbour, which is a muddy environment. However, Spirobranchus (as Pomatoceros) triqueter has been noted to also occur in areas where there is little or no silt present (Price et al., 1980).

Barnes and Bagenal (1951) found that growth rate of Balanus crenatus epizoic on Nephrops norvegicus was considerably slower than animals on raft exposed panels. This was attributed to reduced currents and increased silt loading of water in the immediate vicinity of Nephrops norvegicus. In dredge disposal areas in the Weser estuary, Germany, where turbidity is 35% above the natural rate of 10-100 mg/l, the abundance of Balanus crenatus was lower than in reference areas (Witt et al., 2004). Separating the effect of increased suspended solids from increased sedimentation and changes in sediment from sediment dumping is problematic, however (Witt et al., 2004). Balanids may stop filtration after silt layers of a few millimetres have been discharged (Witt et al., 2004), as the feeding apparatus is very close to the sediment surface.

A significant decrease in suspended organic particles may reduce food input to the biotope resulting in reduced growth and fecundity of barnacles and Spirobranchus triqueter. However, local primary productivity may be enhanced where suspended sediments decrease, increasing food supply. Decreased suspended sediment may increase macroalgal competition enhancing diversity but is considered unlikely to significantly change the character of the biotope as colonisation by larger brown macroalgae is limited by the friability of the surface which is unsuitable for attachment.

Gyory et al., (2013) found that increased turbidity triggered the release of larvae by Semibalanus balanoides, a response which may allow larval release to be timed with high levels of phytoplankton and at times where predation on larvae may be lowered due to the concentration of particles. Storm events that stir up sediments are also associated with larval release (Gyory & Pineda, 2011).

Sensitivity assessment. Overall biotope resistance is assessed as ‘High’ to an increase in suspended solids. Resilience is categorised as ‘High’ (by default). The biotope is considered to be ’Not sensitive’ to decreased suspended solids.

As small, sessile species attached to the substratum, siltation at the pressure benchmark would bury spirorbids, barnacles and Spirobranchus triqueter. Holme and Wilson (1985) described a Pomatoceros-Balanus assemblage on ‘hard surfaces subjected to periodic sever scour and ‘deep submergence by sand or gravel’ in the English Channel. They inferred that the Pomatoceros-Balanus assemblage was restricted to fast-growing settlers able to establish themselves in short periods of stability during summer months (Holme and Wilson 1985), as all fauna were removed in the winter months. Barnacles may stop filtration after silt layers of a few millimetres have been discharged as the feeding apparatus is very close to the sediment surface (Witt et al., 2004). In dredge disposal areas in the Weser estuary, Germany, where the modelled exposure to sedimentation was 10mm for 25 days, with the centre of the disposal ground exposed to 65 mm for several hours before dispersal, Balanus crenatus declined in abundance compared to reference areas. (Witt etal., 2004). However, separating the effect of sedimentation from increased suspended solids and changes in sediment from sediment dumping was problematic (Witt et al., 2004).

In a review of the effects of sedimentation on rocky coast assemblages, Airoldi (2003) outlined the evidence for the sensitivity of encrusting coralline algae to sedimentation. The reported results are contradictory with some authors suggesting that coralline algae are negatively affected by sediments while others report that encrusting corallines are often abundant or even dominant in a variety of sediment impacted habitats (Airoldi, 2003 and references therein). Crustose corallines have been reported to survive under a turf of filamentous algae and sediment for 58 days (the duration of experiment) in the Galapagos (species not identified, Kendrick, 1991). The crustose coralline Hydrolithon reinboldii, has also been reported to survive deposition of silty sediments on subtidal reefs off Hawaii (Littler, 1973). In an experimental study, Balata et al. (2007) enhanced sedimentation on experimental plots in the Mediterranean (close to Tuscany) by adding 400 g of fine sediment every 45 days on plots of 400 cm2 for 1 year. Nearby sites with higher and lower levels of sedimentation were assessed as control plots. Some clear trends were observed. Crustose corallines declined at medium and high levels of sedimentation (Balata et al., 2007). The experiment relates to chronic low levels of sedimentation rather than a single acute event, as in the pressure benchmark, however the trends observed are considered to have some relevance to the pressure assessment.

Field observations and laboratory experiments have highlighted the sensitivity of limpets to sediment deposition (see also the ‘heavy’ siltation pressure for further information). Airoldi & Hawkins (2007) tested the effects of different grain sizes and deposit thickness in laboratory experiments using Patella vulgata. Sediments were added as a ‘fine’ rain to achieve deposit thicknesses of approximately 1mm, 2 mm, and 4 mm in controlled experiments and grazing and mortality observed over 8-12 days. Limpets were more sensitive to sediments with a higher faction of fines (67% silt) than coarse (58% sand). Coarse sediments of thicknesses approximately 1, 2 and 4 mm decreased grazing activity by 35, 45 and 50 % respectively. At 1 and 2 mm thicknesses, fine sediments decreased grazing by 40 and 77 %. The addition of approximately 4 mm of fine sediment completely inhibited grazing. Limpets tried to escape the sediment but lost attachment and died after a few days (Airoldi & Hawkins, 2007).

Observations on exposed and sheltered shores with patches of sediment around Plymouth in the south west of England ,found that Patella vulgata abundances were higher where deposits were absent. The limpets were locally absent in plots with 50-65% sediment cover (Airoldi & Hawkins, 2007). Littler et al., (1983) found that the another limpet species, Lottia gigantea on southern Californian shores was restricted to refuges from sand burial on shores subject to periodic inundation by sands.

Sensitivity assessment. Based on the presence of the characterizing and associated species in biotopes subject to sedimentation and scour (such as CR.MCR.EcCr.UrtScr), biotope resistance to this pressure, at the benchmark, is assessed as 'High', resilience is assessed as 'High' (by default) and the biotope is considered to be 'Not sensitive'. The assessment considers that sediments are rapidly removed from the biotope and that the scour tolerance of the characterizing animal species and encrusting corallines would prevent significant mortalities although some damage and abrasion may occur. However, if the deposit remained in place; i.e. due to the scale of the pressure or where biotopes were sheltered, or only seasonally subject to water movements or where water flows and wave action were reduced e.g. by the presence of tidal barrages, then resistance would be lower and sensitivity would be greater. Even small deposits of sediments are likely to result in local removal of limpets. The level of impact will depend on the magnitude and duration of impact. Resistance to siltation is assessed as ‘Low’ for Patella vulgata based primarily on observations and experiments of Airoldi & Hawkins, (2007), who demonstrated negative effects at deposit thicknesses far lower than the pressure benchmark. Small patches subject to a single impact may recover rapidly via adult migration. As this species is not considered to characterize the biotope, the lower sensitivity based on the barnacles and spirorbids is presented in the risk assessment table.

This biotope is described as subject to scouring from abrasion by mobile sediments (Connor et al., 2004). The characterizing species occur in biotopes subject to sedimentation and scour (such as CR.MCR.EcCr.UrtScr) and are therefore likely to tolerate intermittent episodes of fine sediment movement and deposition. At the pressure benchmark ‘heavy deposition’ represents a considerable thickness of deposit and complete burial of the characterizing species would occur. Removal of the sediments by wave action and tidal currents would result in considerable scour. The effect of this pressure will be mediated by the length of exposure to the deposit and the nature of the deposit.

As small, sessile species attached to the substratum, siltation at the pressure benchmark would bury barnacles and spirorbids. The lower limits of Semibalanus balanoides (as Balanus balanoides) appear to be set by levels of sand inundation on sand-affected rocky shores in New Hampshire (Daly & Mathieson, 1977. Holme and Wilson (1985) described a Pomatoceros-Balanus assemblage on ‘hard surfaces subjected to periodic sever scour and ‘deep submergence by sand or gravel’ in the English Channel. They inferred that the Pomatoceros-Balanus assemblage was restricted to fast-growing settlers able to establish themselves in short periods of stability during summer months (Holme and Wilson, 1985), as all fauna were removed in the winter months. Barnacles may stop filtration after silt layers of a few millimetres have been discharged as the feeding apparatus is very close to the sediment surface (Witt et al., 2004). In dredge disposal areas in the Weser estuary, Germany, where the modelled exposure to sedimentation was 10mm for 25 days, with the centre of the disposal ground exposed to 65 mm for several hours before dispersal, Balanus crenatus declined in abundance compared to reference areas. (Witt et al., 2004). However, separating the effect of sedimentation from increased suspended solids and changes in sediment from sediment dumping was problematic (Witt et al., 2004).

In a review of the effects of sedimentation on rocky coast assemblages, Airoldi (2003) outlined the evidence for the sensitivity of encrusting coralline algae to sedimentation. The reported results are contradictory with some authors suggesting that coralline algae are negatively affected by sediments while others report that encrusting corallines are often abundant or even dominant in a variety of sediment impacted habitats (Airoldi, 2003 and references therein). Crustose corallines have been reported to survive under a turf of filamentous algae and sediment for 58 days (the duration of experiment) in the Galapagos (species not identified, Kendrick, 1991). The crustose coralline Hydrolithon reinboldii, has also been reported to survive deposition of silty sediments on subtidal reefs off Hawaii (Littler, 1973). In an experimental study, Balata et al. (2007) enhanced sedimentation on experimental plots in the Mediterranean (close to Tuscany) by adding 400 g of fine sediment every 45 days on plots of 400 cm2 for 1 year. Nearby sites with higher and lower levels of sedimentation were assessed as control plots. Some clear trends were observed. Crustose corallines declined at medium and high levels of sedimentation (Balata et al., 2007). The experiment relates to chronic low levels of sedimentation rather than a single acute event, as in the pressure benchmark, however the trends observed are considered to have some relevance to the pressure assessment.

Field observations and laboratory experiments have highlighted the sensitivity of limpets to sediment deposition (see also the ‘heavy’ siltation pressure for further information). Airoldi & Hawkins (2007) tested the effects of different grain sizes and deposit thickness in laboratory experiments using Patella vulgata. Sediments were added as a ‘fine’ rain to achieve deposit thicknesses of approximately 1mm, 2 mm, and 4 mm in controlled experiments and grazing and mortality observed over 8-12 days. Limpets were more sensitive to sediments with a higher faction of fines (67% silt) than coarse (58% sand). Coarse sediments of thicknesses approximately 1, 2 and 4 mm decreased grazing activity by 35, 45 and 50 % respectively. At 1 and 2 mm thicknesses, fine sediments decreased grazing by 40 and 77 %. The addition of approximately 4 mm of fine sediment completely inhibited grazing. Limpets tried to escape the sediment but lost attachment and died after a few days (Airoldi & Hawkins, 2007). Observations on exposed and sheltered shores with patches of sediment around Plymouth in the south west of England , found that Patella vulgata abundances were higher where deposits were absent. The limpets were locally absent in plots with 50-65% sediment cover (Airoldi & Hawkins, 2007). Littler et al., (1983) found that another limpet species, Lottia gigantea on southern Californian shores was restricted to refuges from sand burial on shores subject to periodic inundation by sands.

Sensitivity assessment. Sensitivity to this pressure will be mediated by site-specific hydrodynamic conditions and the footprint of the impact. Where a large area is covered sediments may be shifted by wave and tides within the cave rather than removed. Resistance is assessed as ‘Medium’ as the biotope is exposed to frequent abrasion and scouring (the impact may be mitigated by rapid removal of the deposit) but some removal and mortalities may occur. Resilience is assessed as ‘High’ based on re-growth from the scour-tolerant, surviving bases of the encrusting corallines and larval recolonization by barnacles, spirorbids and Spirobranchus triqueter. Biotope sensitivity is therefore assessed as 'Low'. Even small deposits of sediments are likely to result in local removal of limpets. The level of impact will depend on the magnitude and duration of impact. Resistance to siltation is assessed as ‘Low’ for Patella vulgata based primarily on observations and experiments of Airoldi & Hawkins, (2007), who demonstrated negative effects at deposit thicknesses far lower than the pressure benchmark. Small patches subject to a single impact may recover rapidly via adult migration. As this species is not considered o characterize the biotope, the lower sensitivity based on the barnacles and spirorbids is presented in the risk assessment table.

Not assessed. Thompson et al., (2004) demonstrated that Semibalanus balanoides, kept in aquaria, ingested microplastics within a few days. However, the effects of the microplastics on the health of exposed individuals have not been identified. There is currently no evidence to assess the level of impact. Not relevant. Wave action on exposed shores is likely to generate high levels of underwater noise. Other sources are not considered likely to result in effects on the biotope.

Light penetration is a key factor structuring the cave biotope. Encrusting corallines and other shade-tolerant algae grow closer to the entrance where light availability allows. Encrusting corallines can occur in deeper water than other algae where light penetration is limited. Samples of Lithophyllum impressum suspended from a raft and shaded (50-75% light reduction) continued to grow over two years (Dethier, 1994). An increase in light in the spectrum that supports photosynthesis may allow algae including Rhodochorton purpureum and Pilinia maritima which are found within caves (Connor et al., 2004) to colonise more surface area, altering the structure of the biotope. Increased light may also reduce recruitment of spirorbids which have been found to recruit preferentially to shaded areas (Saunders & Connell, 2001).

The animal species are not light dependent. Spirobranchus triqueter is found in a variety of light environments from shallow sublittoral biotopes where light levels are relatively high, to deeper sites that are aphotic (De Kluijver, 1993).

Balanus crenatus possesses a rudimentary eye and can detect and respond to sudden shading which may be an anti-predator defence (Forbes et al., 1971). Balanus crenatus tend to orient themselves when settling, with the least light sensitive area directed towards the light (Forbes et al., 1971), so that the more sensitive area can detect shading from predator movements in the area where light availability is lower (Forbes et al., 1971).

Semibalanus balanides sheltered from the sun grew bigger than unshaded individuals (Hatton, 1938; cited in Wethey, 1984), although the effect may be due to indirect cooling effects rather than shading. Barnacles are also frquently found under algal canopies suggesting that they are tolerant of shading. Light levels have also been demonstrated to influence a number of phases of the reproductive cycle in Semibalanus balanoides. In general light inhibits aspects of the breeding cycle. Penis development is inhibited by light (Barnes & Stone, 1972) while Tighe-Ford (1967) showed that constant light inhibited gonad maturation and fertilization. Davenport & Crisp (unpublished data from Menai Bridge, Wales, cited from Davenport et al., 2005) found that experimental exposure to either constant darkness, or 6 h light: 18 h dark photoperiods induced autumn breeding in Semibalanus. They also confirmed that very low continuous light intensities (little more than starlight) inhibited breeding. Latitudinal variations in timing of the onset of reproductive phases (egg mass hardening) have been linked to the length of darkness (night) experienced by individuals rather than temperature (Davenport et al., 2005). Changes in light levels associated with climate change (increased cloud cover) were considered to have the potential to alter timing of reproduction (Davenport et al., 2005) and to shift the range limits of this species southward. However, it is not clear how these findings may reflect changes in light levels from artificial sources, and whether observable changes would occur at the population level as a result. There is, therefore, 'No evidence' on which to base an assessment.

Sensitivity assessment. The key characterizing animal species colonize a broad range of light environments, from intertidal to deeper sub tidal and shaded understorey habitats and are considered to be unaffected by increased shade or more light penetration. However an increase in light in the spectrum that supports photosynthesis may increase algal growth altering the character of the biotope. Some specialist cave species may colonize depending on the presence of source populations. The biotope is therefore considered to have ‘Low’ resistance and ‘High’ resilience following restoration of typical conditions (as crustose corallines and characterizing animals are likely to survive). Sensitivity is therefore considered to be ‘Low’.

No direct evidence was found to assess this pressure. Barriers that reduce the degree of tidal excursion may alter larval supply to suitable habitats from source populations. Conversely the presence of barriers may enhance local population supply by preventing the loss of larvae from enclosed habitats. As the larvae of Balanus crenatus and Semibalanus balanoides and other species such as Patella vulgata are planktonic and are transported by water movements, barriers that reduce the degree of tidal excursion may alter larval supply to suitable habitats from source populations. However the presence of barriers may enhance local population supply by preventing the loss of larvae from enclosed habitats. Barriers and changes in tidal excursion are not considered relevant to the characterizing crusting corallines as species dispersal is limited by the rapid rate of settlement and vegetative growth from bases rather than reliance on recruitment from outside of populations. Resistance to this pressure is assessed as 'High' and resilience as 'High' by default. This biotope is therefore considered to be 'Not sensitive'.

Many of the animal species within the biotope probably respond to light levels, detecting shade and shadow to avoid predators, and day length in their behavioural or reproductive strategies. However, their visual acuity is probably very limited and they are unlikely to respond to visual disturbance at the benchmark level. This pressure is, therefore, assessed as ‘Not relevant’.

Balanus crenatus possesses a rudimentary eye and can detect and respond to sudden shading which may be an anti-predator defence (Forbes et al., 1971). Balanus crenatus tend to orient themselves when settling, with the least light sensitive area directed towards the light (Forbes et al., 1971), so that the more sensitive area can detect shading from predator movements in the area where light availability is lower (Forbes et al., 1971).

The high levels of scour in this biotope will limit establishment of all but the most scour resistant invasive non-indigenous species (INIS) and no direct evidence was found for effects of INIS on this biotope. The low levels of light within this biotope, particularly the rear walls of caves, are considered to also inhibit invasive algal species.

The Australasian barnacle Austrominius (previously Elminius) modestus was introduced to British waters on ships during the second world war. Increased warming has allowed the Australian barnacle Austrominius (formerly, Elminiusi) modestus, to dominate sites previously occupied by Semibalanus balanoides and Balanus crenatus (Witte, 2010). However, on settlement panels deployed in SW Ireland, Austrominius modestus initially dominated panels in the lower subtidal but post-recruitment mortality over a year allowed Balanus crenatus to become the dominant barnacle (Watson et al., 2005). Balanus crenatus and Austrominius modestus have shown recruitment differences which may alter the seasonal dominance patterns (Witte et al., 2010). In generalits overall effect on the dynamics of rocky shores has been small as Austrominius modestus has simply replaced some individuals of a group of co-occurring barnacles (Raffaelli & Hawkins, 1999). Although present, monitoring indicates it has not outnumbered native barnacles in the Isle of Cumbrae (Gallagher et al., 2015), it may dominate in estuaries where it is more tolerant of lower salinities than Semibalanus balanoides (Gomes-Filho, et al., 2010).

Two non-native spirobids – Dexiospira oshoroensis and Pileolaria rosepigmentata - were found on the non-native algae Sargassum muticum in Portsmouth (Knight-Jones et al., 1975). Invasive tubeworms are reported from UK harbours (Thorp et al., 1986) and are likely to be well established in areas with large volumes of ship traffic.

The tunicates Didemnum vexillum and Asterocarpa humilis, the hydroid Schizoporella japonica and the bryozoan Watersipora subatra (Bishop, 2012c, Bishop, 2015a and b; Wood, 2015) are currently only recorded from artificial hard substratum in the UK and it is not clear what their established range and impacts in the UK would be. Didemnum vexillum occurs in tide pools in other areas where it has become established (Bishop, 2012c) and can have substantial effects on communities, similarly the tunicates Corella eumyota and Botrylloides violaceus can smother rock habitats (Bishop, 2011b and 2012b).

Sensitivity assessment. Overall, there is little evidence of this biotope being adversely affected by non-native species and scouring of this biotope by mobile sediments is likely to limit establishment of all but robust species. Replacement of native barnacles and spirorbids by non-natives alters the identity of the species present but has little impact on biotope character and function. Resistance is therefore assessed as 'High', and resilience as 'High' (by default), and the biotope is considered to be 'Not sensitive'.

No evidence was found that microbial pathogens cause high levels of disease or mortality in this biotope.

The cherizing species Semibalanus balanoides are considered subject to persistent, low levels of infection by pathogens and parasites. Barnacles are parasitised by a variety of organisms and, in particular, the cryptoniscid isopod Hemioniscus balani, in which heavy infestation can cause castration of the barnacle. At usual levels of infestation these are not considered to lead to high levels of mortality. The associated species Patella vulgata has been reported to be infected by the protozoan Urceolaria patellae (Brouardel, 1948) at sites sheltered from extreme wave action in Orkney. Baxter (1984) found shells to be infested with two boring organisms, the polychaete Polydora ciliata and a siliceous sponge Cliona celata.

Diseased encrusting corallines were first observed in the tropics in the early 1990’s when the bacterial pathogen Coralline Lethal Orange Disease (CLOD) was discovered (Littler & Littler, 1995). All species of articulated and crustose species tested to date are easily infected by CLOD and it has been increasing in occurrence at sites where first observed and spreading through the tropics. Another bacterial pathogen causing a similar CLOD disease has been observed with a greater distribution and a black fungal pathogen first discovered in American Samoa has been dispersing (Littler & Littler, 1998). An unknown pathogen has also been reported to lead to white ‘target-shaped’ marks on corallines, again in the tropic (Littler et al., 2007). No evidence was found that these pathogens are impacting temperate coralline habitats.

Sensitivity assessment. Based on the lack of reported mortalities of the characterizing species and the available evidence for the coralline crust, the biotope is judged to have ‘High’ resistance to this pressure. By default resilience is assessed as ‘High’ and the biotope is classed as ‘Not sensitive’ at the pressure benchmark.

Direct, physical impacts from harvesting are assessed through the abrasion and penetration of the seabed pressures. The sensitivity assessment for this pressure considers any biological/ecological effects resulting from the removal of target species on this biotope. Limpets may be gathered recreationally for consumption but removal of this species is not considered to alter the character of the biotope through its loss as shade and scour, rather than grazing, are the key factors limiting the presence of algae (Connor et al., 2004). No commercial application or harvesting of other characterizing or associated species is described in the literature and this pressure is therefore considered to be 'Not relevant'.

Incidental removal of the key characterizing species would alter the character of the biotope, resulting in reclassification and the loss of species richness. The ecological services such as primary and secondary production, provided by characterizing and associated species, would also be lost. As most species present in this biotope are relatively large, conspicuous and either sedentary or attached to rock surfaces that have little protection against removal.

Sensitivity assessment. Removal of a large percentage of the characterizing species resulting in bare rock would alter the character of the biotope, species richness and ecosystem function. Resistance is therefore assessed as ‘Low’ and recovery as ‘High’, so that biotope sensitivity is assessed as 'Low’.

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